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RESEARCH ARTICLE
Contents Vol 22(2)

Molecular and morphological systematics of hypogean schizomids (Schizomida : Hubbardiidae) in semiarid Australia

Mark S. Harvey A D , Oliver Berry B , Karen L. Edward A and Garth Humphreys C
+ Author Affiliations
- Author Affiliations

A Department of Terrestrial Invertebrates, Western Australian Museum, Locked Bag 49, Welshpool DC, Western Australia 6986, Australia.

B School of Animal Biology, The University of Western Australia, Crawley, Western Australia 6009, Australia.

C Biota Environmental Sciences Pty Ltd, PO Box 155, Leederville, Western Australia 6903, Australia.

D Corresponding author. Email: mark.harvey@museum.wa.gov.au

Invertebrate Systematics 22(2) 167-194 https://doi.org/10.1071/IS07026
Submitted: 11 June 2007  Accepted: 30 January 2008   Published: 12 May 2008

Abstract

We used molecular and morphological techniques to study troglobitic schizomids inhabiting a variety of subterranean landforms in semiarid Western Australia. The study was designed to explore the taxonomic and phylogenetic status of newly discovered populations of subterranean schizomids. Molecular sequences of the mitochondrial DNA (mtDNA) cytochrome oxidase subunit 1 (COI) and small subunit rRNA (12S) were obtained from a total of 73 schizomid specimens. Populations sampled from boreholes within mesa landforms in the Robe Valley were highly genetically distinct from species of Draculoides Harvey, 1992 found elsewhere in the Pilbara (Cape Range and Barrow Island). Pronounced genetic structuring was also evident at a fine spatial scale within the Robe Valley, with populations from each of the mesas examined exhibiting unique and highly divergent mtDNA lineages. These molecular data were generally supported by small but significant morphological features, usually in the secondary male structures, but some species were represented only by female specimens that possessed more conservative morphologies. The molecular data defined two major in-group clades, which were supported by morphological differences. One clade was widespread and included the type species of Draculoides, D. vinei (Harvey), along with D. bramstokeri Harvey & Humphreys, D. brooksi Harvey, D. julianneae Harvey, D. mesozeirus, sp. nov. and D. neoanthropus, sp. nov. The second clade was restricted to the Robe Valley and deemed to represent a new genus, Paradraculoides, which included four new species P. anachoretus, sp. nov., P. bythius, sp. nov., P. gnophicola, sp. nov. and P. kryptus, sp. nov. (type species).


Acknowledgements

This study was funded by Robe River Iron Associates and Chevron Australia Pty Ltd. We would like to thank Michael Greenham, Dan Kamien, Lee Mould, Tim Moulds, Bill Humphreys, Darren Brooks, Michael Rix, Robert Raven and Jay Anderson for supplying the schizomid specimens used in the analysis, and Julianne Waldock for laboratory assistance. We also thank Lorenzo Prendini, Cor Vink and Gonzalo Giribet for providing useful advice on PCR primers, and Peter Landman and Fiona Bell of RRIA for facilitating the project. We also thank Lorenzo Prendini, Camilla Myers and an anonymous referee for their helpful comments on a draft of the manuscript.


References


Adams M., Humphreys W. F. (1993) Patterns of genetic diversity within selected subterranean fauna of the Cape Range Peninsula, Western Australia: systematic and biogeographic implications. Records of the Western Australian Museum Supplement 45, 145–164. open url image1

de Armas L. F. (1989) Adiciones al orden Schizomida (Arachnida) en Cuba. Poeyana 387, 1–45. open url image1

de Armas L. F. (2002a) Dos géneros nuevos de Hubbardiidae (Arachnida: Schizomida) de Cuba. Revista Ibérica de Aracnología 5, 3–9. open url image1

de Armas L. F. (2002b) Nuevas especies de Rowlandius Reddell & Cokendolpher, 1995 (Schizomida: Hubbardiidae) de Cuba. Revista Ibérica de Aracnología 6, 149–167. open url image1

de Armas L. F. (2004) Nueva especie de Rowlandius Reddell & Cokendolpher, 1995 (Schizomida: Hubbardiidae) de Cuba oriental. Revista Ibérica de Aracnología 10, 211–214. open url image1

de Armas L. F. (2005) Los esquizómidos troglobios de Cuba (Arachnida: Schizomida). Espelunca 1, 2–5. open url image1

de Armas L.F.,, Teruel R. (2002) Un género nuevo de Hubbardiidae (Arachnida: Schizomida) de las Antilles Mayores. Revista Ibérica de Aracnología 6, 45–52. open url image1

BMR Palaeogeographic Group (1990). ‘Australia: Evolution of a Continent.’ (Australian Bureau of Natural Resources: Canberra, Australia.)

Bond J. E., Sierwald P. (2003) Molecular taxonomy of the Anadenobolus excisus (Diplopoda: Spirobolida: Rhinocricidae) species-group on the Caribbean island of Jamaica. Invertebrate Systematics 17, 515–528.
| Crossref | GoogleScholarGoogle Scholar | open url image1

Bradbury J. H. (2000) Western Australian stygobiont amphipods (Crustacea: Paramelitidae) from the Mt Newman and Millstream regions. Records of the Western Australian Museum 60, 1–102. open url image1

Bryant H. N. (1992) Why autapomorphies should be removed. A reply to Yeates. Cladistics 10, 177–181. open url image1

Clary D. O., Wolstenholme D. R. (1985) The mitochondrial DNA molecule of Drosophila yakuba: Nucleotide sequence, gene organization and genetic code. Journal of Molecular Evolution 22, 252–271.
| Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Cokendolpher J. C., Sissom W. D., Bastawade D. B. (1988) A new Schizomus from the Indian state of Maharashtra, with additional comments on eyed schizomids (Arachnida: Schizomidae). Insecta Mundi 2, 90–96. open url image1

Eberhard S. M., Halse S. A., Humphreys W. F. (2005) Stygofauna in the Pilbara region, north-west Western Australia: a review. Journal of the Royal Society of Western Australia 88, 167–176. open url image1

Edward K. L., Harvey M. S. (2008) Short-range endemism in hypogean environments: the pseudoscorpion genera Tyrannochthonius and Lagynochthonius (Pseudoscorpiones: Chthoniidae) in the semiarid zone of Western Australia. Invertebrate Systematics 22, 259–293.
| Crossref | GoogleScholarGoogle Scholar | open url image1

Farris J. S., Albert V. A., Källersjö M., Lipscomb D., Kluge A. G. (1996) Parsimony jackknifing outperforms neighbor-joining. Cladistics 12, 99–124.
| Crossref | GoogleScholarGoogle Scholar | open url image1

Georgescu M. (1994). Schizomida. In ‘Encyclopaedia Biospeologica. Vol. 1’. (Eds C. Juberthie and V. Decu.) pp. 237–240. (Société de Biospéologie: Moulis and Bucarest.)

Goloboff P. A. (1993). NONA, Version 2.0. (Published by the author: Tucumán, Argentina.)

Harvey M. S. (1988) A new troglobitic schizomid from Cape Range, Western Australia (Chelicerata: Schizomida). Records of the Western Australian Museum 14, 15–20. open url image1

Harvey M. S. (1992) The Schizomida (Chelicerata) of Australia. Invertebrate Taxonomy 6, 77–129.
| Crossref | GoogleScholarGoogle Scholar | open url image1

Harvey M. S. (2000a) Brignolizomus and Attenuizomus, new schizomid genera from Australia (Arachnida: Schizomida: Hubbardiidae). Memorie della Società Entomologica Italiana. Supplemento 78, 329–338. open url image1

Harvey M. S. (2000b) A review of the Australian schizomid genus Notozomus (Hubbardiidae). Memoirs of the Queensland Museum 46, 161–174. open url image1

Harvey M. S. (2001a) The Schizomida (Arachnida) of the Seychelle Islands. Invertebrate Taxonomy 15, 681–693.
| Crossref | GoogleScholarGoogle Scholar | open url image1

Harvey M. S. (2001b) New cave-dwelling schizomids (Schizomida: Hubbardiidae) from Australia. Records of the Western Australian Museum Supplement 64 , 171–185. open url image1

Harvey M. S. (2002) Short-range endemism in the Australian fauna: some examples from non-marine environments. Invertebrate Systematics 16, 555–570.
| Crossref | GoogleScholarGoogle Scholar | open url image1

Harvey M. S. (2003). ‘Catalogue of the Smaller Arachnid Orders of the World: Amblypygi, Uropygi, Schizomida, Palpigradi, Ricinulei and Solifugae.’ (CSIRO Publishing: Melbourne, Australia.)

Harvey M. S., Edward K. L. (2007a) A review of the pseudoscorpion genus Ideoblothrus (Pseudoscorpiones, Syarinidae) from western and northern Australia. Journal of Natural History 41, 445–472.
| Crossref | GoogleScholarGoogle Scholar | open url image1

Harvey M.S., Edward K.L. (2007b) Three new species of cavernicolous goblin spiders (Araneae, Oonopidae) from Australia. Records of the Western Australian Museum 24, 9–17. open url image1

Harvey M. S., Gray M. R., Hunt G. S., Lee D. C. (1993) The cavernicolous Arachnida and Myriapoda of Cape Range, Western Australia. Records of the Western Australian Museum Supplement 45 , 129–144. open url image1

Harvey M. S., Humphreys W. F. (1995) Notes on the genus Draculoides Harvey (Schizomida: Hubbardiidae), with the description of a new troglobitic species. Records of the Western Australian Museum Supplement 52 , 183–189. open url image1

Harvey M. S., Shear W. A., and Hoch H. (2000). Onychophora, Arachnida, myriapods and Insecta. In ‘Subterranean Ecosystems’. (Eds H. Wilkens, D. C. Culver and W. F. Humphreys.) pp. 79–94. (Elsevier: Amsterdam, The Netherlands.)

Howarth F. G. (1988). Environmental ecology of north Queensland caves: or why are there so many troglobites in Australia? In ‘Australian Speleological Federation Tropicon Conference, Lake Tinaroo, Far North Queensland’. (Ed. L. Pearson.) pp. 76–84. (Australian Speleological Federation: Cairns, Australia.)

Humphreys W. F. (1990) The biology of a troglobitic schizomid (Chelicerata: Arachnida) from caves in the semi-arid Cape Range, Western Australia. Acta Zoologica Fennica 190, 181–186. open url image1

Humphreys W. F. (1991) Experimental re-establishment of pulse-driven populations in a terrestrial troglobite community. Journal of Animal Ecology 60, 609–623.
| Crossref | GoogleScholarGoogle Scholar | open url image1

Humphreys W. F. (1993a) The significance of the subterranean fauna in biogeographical reconstruction: examples from Cape Range Peninsula, Western Australia. Records of the Western Australian Museum Supplement 45 , 165–192. open url image1

Humphreys W. F. (1993b) Cave fauna in semi-arid tropical Western Australia: a diverse relict wet-forest litter fauna. Memoires de Biospeologie 20, 105–110. open url image1

Humphreys W. F. (1998) Phaconeura (Homoptera: Meenoplidae) attended by ants of the genus Paratrechina (Hymenoptera: Formicidae) in caves. Australian Entomologist 25, 23–27. open url image1

Humphreys W. F. (1999). Relict stygofaunas living in sea salt, karst and calcrete habitats in arid northwestern Australia contain many ancient lineages. In ‘The Other 99%. The Conservation and Biodiversity of Invertebrates’. (Eds W. Ponder and D. Lunney.) pp. 219–227. (Royal Zoological Society of New South Wales: Mosman, Australia.)

Humphreys W. F. (2000). The hypogean fauna of the Cape Range Peninsula and Barrow Island, northwestern Australia. In ‘Subterranean Ecosystems’. (Eds H. Wilkens, D. C. Culver and W. F. Humphreys.) pp. 581–601. (Elsevier: Amsterdam, The Netherlands.)

Humphreys W. F. (2001a) Groundwater calcrete aquifers in the Australian arid zone: the context to an unfolding plethora of stygal biodiversity. Records of the Western Australian Museum Supplement 64 , 63–83. open url image1

Humphreys W. F. (2001b) The subterranean fauna of Barrow Island, northwestern Australia, and its environment. Memoires de Biospeologie 28, 107–127. open url image1

Humphreys W. F. (2006) Aquifers: the ultimate groundwater-dependent ecosystems. Australian Journal of Botany 54, 115–132.
| Crossref | GoogleScholarGoogle Scholar | open url image1

Humphreys W. F., Collis G. (1990) Water loss and respiration of cave arthropods from Cape Range, Western Australia. Comparative Biochemistry and Physiology. A: Comparative Physiology 95, 101–107.
| Crossref | GoogleScholarGoogle Scholar | open url image1

Humphreys W. F., Adams M., Vine B. (1989) The biology of Schizomus vinei (Chelicerata: Schizomida) in the caves of Cape Range, Western Australia. Journal of Zoology 217, 177–201. open url image1

Kambhampati S., Smith P. T. (1995) PCR primers for the amplification of four insect mitochondrial gene fragments. Insect Molecular Biology 4, 233–236.
| Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Kocher T. D., Thomas W. K., Meyer A., Edwards S. V., Pääbo S., Villablanca F. X., Wilson A. C. (1989) Dynamics of mitochondrial DNA evolution in animals: amplification and sequencing with conserved primers. Proceedings of the National Academy of Sciences of the United States of America 86, 6196–6200.
| Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

MacPhail M. K., Stone M. S. (2004) Age and palaeoenvironmental constraints on the genesis of the Yandi channel iron deposits, Marillana Formation, Pilbara, northwestern Australia. Australian Journal of Earth Sciences 51, 497–520.
| Crossref | GoogleScholarGoogle Scholar | open url image1

Maddison W. P., and Maddison D. R. (2005). Mesquite: A modular system for evolutionary analysis. Version 2.0. Available at http://mesquiteproject.org [Verified March 2008].

Main B. Y. (1980) Record of the arachnid order Schizomida in Western Australia. Western Australian Naturalist 14, 236–237. open url image1

Martín J. L., Oromí P. (1984) Consideraciones sobre la presencia de Schizomus portoricensis Chamberlin, 1922) (Arach. Schizomida) en cuevas de Tenerife (Islas Canarias). Boletín de la Sociedad Entomológica de España 8, 265–270. open url image1

Morton S. R., Short J., Barker R. D. (1995) Refugia for biological diversity in arid and semi-arid Australia. Biodiversity Series, Biodiversity Unit, Department of the Environment. Sport and Territories 4, 1–171. open url image1

Nixon K. C. (2002). Winclada. Version 1.00.08. (Published by the author. Ithaca: New York, USA.) Available at http://www.cladistics.com/ [Verified March 2008].

Nixon K. C., Wheeler Q. D. (1990) An amplification of the phylogenetic species concept. Cladistics 6, 211–223. open url image1

Page R. D. M. (2001). NEXUS Data Editor for Windows, version 0.5.0. Available from http://taxonomy.zoology.gla.ac.uk/rod/NDE/nde.html [Verified April 2008].

Paquin P., Hedin M. C. (2004) The power and perils of ‘molecular taxomomy’: a case study of eyeless and endangered Cicurina (Araneae: Dictynidae) from Texas caves. Molecular Ecology 13, 3239–3255.
| Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Peck S. B. (1980) Climatic change and the evolution of cave invertebrates in the Grand Canyon. Bulletin of the National Speleological Society [not seen.] 42, 53–60. open url image1

Posada D., Crandall K. A. (1998) Modeltest: testing the model of DNA substitution. Bioinformatics 14, 817–818.
| Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Prendini L. (2000) Phylogeny and classification of the superfamily Scorpionoidea Latreille 1802 (Chelicerata, Scorpiones): an exemplar approach. Cladistics 16, 1–78.
| Crossref | GoogleScholarGoogle Scholar | open url image1

Ramanaidou E. R., Morris R. C., Horwitz R. C. (2003) Channel iron deposits of the Hamersley Province, Western Australia. Australian Journal of Earth Sciences 50, 669–690.
| Crossref | GoogleScholarGoogle Scholar | open url image1

Reddell J. R., Cokendolpher J. C. (1991) Redescription of Schizomus crassicaudatus (Pickard-Cambridge) and diagnoses of Hubbardia Cook, Stenochrus Chamberlin, and Sotanostenochrus new genus, with description of a new species of Hubbardia from California (Arachnida: Schizomida: Hubbardiidae). Pearce Sellards Series 47, 1–24. open url image1

Reddell J. R., Cokendolpher J. C. (1995) Catalogue, bibliography, and generic revision of the order Schizomida (Arachnida). Texas Memorial Museum. Speleological Monographs 4, 1–170. open url image1

Rowland J. M., Reddell J. R. (1979a) The order Schizomida (Arachnida) in the New World. I. Protoschizomidae and dumitrescoae group (Schizomidae: Schizomus). The Journal of Arachnology 6, 161–196. open url image1

Rowland J. M., Reddell J. R. (1979b) The order Schizomida (Arachnida) in the New World. II. Simonis and brasiliensis groups (Schizomidae: Schizomus). The Journal of Arachnology 7, 89–119. open url image1

Rowland J. M., Reddell J. R. (1980) The order Schizomida (Arachnida) in the New World. III. Mexicanus and pecki groups (Schizomidae: Schizomus). The Journal of Arachnology 8, 1–34. open url image1

Rowland J. M., Reddell J. R. (1981) The order Schizomida (Arachnida) in the New World. IV. Goonightorum and briggsi groups (Schizomidae: Schizomus). The Journal of Arachnology 9, 19–46. open url image1

Sissom W. D. (1980) The eyed schizomids, with a description of a new species from Sumatra (Schizomida: Schizomidae). The Journal of Arachnology 8, 187–192. open url image1

Swofford D. L. (1998). ‘PAUP*: Phylogenetic Analysis Using Parsimony (and other methods).’ (Sinauer Associates: Sunderland, MA, USA.)

Teruel R. (2000) Una nueva especie de Rowlandius Reddell & Cokendolpher, 1995 (Schizomida: Hubbardiidae) de Cuba Oriental. Revista Ibérica de Aracnología 1, 45–47. open url image1

Teruel R. (2003) Adiciones a la fauna cubana de esquizómidos, con la descripción de un nuevo género y nueve especies nuevas de Hubbardiidae (Arachnida: Schizomida). Revista Ibérica de Aracnología 7, 39–69. open url image1

Teruel R. (2004) Nuevas adiciones a la fauna de esquizómidos de Cuba oriental, con la descripción de cuatro nuevas especies (Schizomida: Hubbardiidae). Revista Ibérica de Aracnología 9, 31–42. open url image1

Teruel R., de Armas L. F. (2002) Un género nuevo de Hubbardiidae (Arachnida: Schizomida) del occidente de Cuba. Revista Ibérica de Aracnología 6, 91–94. open url image1

Thackway R., and Cresswell I. D. (Eds) (1995). ‘An Interim Biogeographic Regionalisation for Australia: A Framework for Establishing the National System of Reserves, Version 4.0.’ (Australian Nature Conservation Agency: Canberra, Australia.)

Thompson J., Higgins D., Gibson T. (1994) CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Research 22, 4673–4680.
| Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Truswell E. M. (1990). Australian rainforests: the 100 million year record. In ‘Australian Tropical Rainforests’. pp. 1–22. (CSIRO: Melbourne, Australia.)

Twidale C. R. (2003) The enigma of survival: problems posed by very old paleosurfaces. Physical Geography 24, 26–60.
| Crossref | GoogleScholarGoogle Scholar | open url image1

Vine B., Knott B., Humphreys W. F. (1988) Observations on the environment and biology of Schizomus vinei (Chelicerata: Schizomida) from Cape Range, Western Australia. Records of the Western Australian Museum 14, 21–34. open url image1

Yeates D. K. (1992) Why remove autapomorphies? Cladistics 8, 387–389.
| Crossref | GoogleScholarGoogle Scholar | open url image1